upper jurassic pleurotomariidae (gastropoda) from … · 2019. 12. 14. · jurassic) age....

THE NAUTILUS 121(2):76-89, 2007 Page 76

Upper Jurassic Pleurotomariidae (Gastropoda) from southwestern Madagascar

M. G. Harasewych Department of Invertebrate Zoolog)' National Museum of Natural History Smithsonian Institution P.O. Box 37012 Washington, DC 20013-7012 USA [email protected]

Steffen Kiel Earth Sciences University of Leeds Leeds LS2 9JT UNITED KINGDOM and Department of Paleobiology National Museum of Natural History Smithsonian Institution Washington, DC 20013-7012 USA [email protected]

ABSTRACT

This paper describes four new species of Upper Jurassic Pleu- rotomariidae from southwestern Madagascar: Obornella thompsonorum, Bathrotomaria annejojfeae, Bathrotomaria bedetteae, and Leptomaria takahashii. In addition, the previously described Leptomaria texta Delpey, 1948, is reassigned to the genus Ohoimella. Comparison of this fauna wdth that of the geographically proximal Kutsch region of northwestern India reveals it to consist of representati\'es of wide-ranging Tethyan genera, but also to exhibit strong endemism at the species level.

evolved an increasing endemism within the Indo- Madagascan fauna! province that was formed as the Tethys Ocean widened between Laurasia and eastern Gondwana, and a seaway started to develop between East and West Gondwana in the latest Jurassic (Titho- nian) (Hay et al, 1999; Shome et al., 2004).

The scope of the present study is to review the Oxfor- dian (Upper Jurassic) pleurotomariid fauna of Madagas- car, describe ñve species, four of them new, and to review the relationships and biogeography of these pleurotomariids.

INTRODUCTION GEOLOGICAL SETTING

A substantial number of well preserved Upper Jurassic pleurotomariid gastropods have recently been uncovered as a byproduct of commercial mining for ammonites in southwestern Madagascar. Six specimens, representing four species, were kindly made available to us for study by Mr. Chris Takahashi. The pleurotomariids and ammonites were dug by \'illagers from pit quarries near the town of Zakaraha, in southwestern Madagascar.

A survey of the literature on the Mesozoic gastropod fauna of Madagascar (e.g., Delpey, 1948; CoUignon, 1949: CoUignon, 1959; Kiel, 2006) revealed the majority of pleurotomariid species known from Madagascar to be of Cretaceous age, with only a single species, Leptomaria texta Delpey, 1948, reported from Jurassic strata. Al- though the sample available to us is of limited size and stratigraphie range, it expands our insight into the Juras- sic pleurotomariid fauna of Madagascar. Like the well documented Jurassic pleurotomariid fauna of the Kutsch (also spelled Kachchh) region of western India (Jaitley et al., 2000; Das, 2002; Das et al, 2005), the Madagascar fauna reveals Tethyan affinities at the generic level, yet exhibits endemism at the species level. Both faunas

Rifting between Africa and Madagascar produced three large sedimentary basins along the west coast of Mada- gascar. These are, from North to South, the Ambilobe (or Diego), Mahajanga (also spelled Majunga), and Moron- dava basins. Sedimentation in this region commenced in the Carboniferous with the deposition of the Gondwanan Karoo sequences and equivalents. The first marine deposits resulting from the break-up of the Gondwana su- percontinent are of Toarcian (late Lower Jurassic) age. From then on, alternating shallow marine, brackish, and fluviatile sediments were deposited in these basins. The pleurotomariids described here are from the Morondava basin, the southernmost of the three basins. Bio- and lithostratigraphic work in this basin is difficult because outcrops are few, index fossils are often not available, and measurable sections are usually short and difficult to correlate with each other. Consequently, only few lithostratigraphic units have been given formation names so far (Besairie and CoUignon, 1972; Luger et al., 1994; Geiger and Schweigert, 2006).

The quarries that produced the specimens described here are located to the west of the town Zakaraha (also

M. G. Harasewych and S. Kiel, 2007 Page 77

spelled Sakkara), one of them north of the Fiherenana River, the other to the south of it (Figures 1, 2). The fossiliferous layers are thin and consist of pink-yellow iron-oolithic limestones, and are overlain by grey mudstones (K. Bändel, pers. comm., 2007). The southern locality is very close, if not identical, with the "Amparam- bato section (VIb)" of Geiger and Schweigert (2006: 99) which was characterized by them as "a highly fossiliferous iron-oolitic limestone and sandstone bed with thin mud-stone interlayers." Accordingly, the oolitic limestones contain an abundant ammonite fauna; the over- laying mudstones contain nodosariid Foraminifera and ostracods (Geiger and Schweigert, 2006). The northern quarry is geographically close to the "Middle-Upper Oxfordian Ankilimena section (XI)" of Geiger and Schweigert (2006). This section is characterized by re- current iron-oolitic limestones, which contain ammonites, rhynchonellids, bivalves, belemnites, echinoderms, and wood debris (Geiger and Schweigert, 2006: 103). Although Geiger and Schweigert (2006) did not report

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pleurotomariids or other gastropods from these sections, the remaining fossil content, their geographic position, and their lithology agrees well with that obsen ed at the ammonite quarries visited by K. Bändel (pers. comm., 2007).

The ammonite fauna of these two localities, especially the presence of Dhosaites cf. primus GoUignon, 1959, suggests a 'middle to upper Argovian (early Oxfordian)' age (GoUignon, 1959; H. Keupp, pers. comm., 2004). The term 'Argowan' has been abandoned due to its in- consistent usage, but largely falls within the range of the Oxfordian (Zeiss, 2003). The sediments considered as 'Argo\dan' by Besairie and GoUignon (1972) correlate with those mapped as late Gallo\dan-early Oxfordian by Geiger and Schweigert (2006). Thus, the pleurotomariids described here are most probably of Oxfordian (Upper Jurassic) age.

SUPRASPEGIFIG GLASSIFIGATION WITHIN PLEUROTOMARIIDAE

The number of genera and subgenera currently recognized within the family Pleurotomariidae (Appendix 1) has nearly doubled since the family was re\iewed in the Treatise of Invertebrate Zoology (Knight et al., 1960). Of the 21 genera and subgenera now recognized (Figure 3), 16 are Mesozoic. Of these, ñve are restricted to the Tri- assic, three to the Jurassic, and only a single subgenus to the Gretaceous. At present, Leptomaria and Conoto- maria are the only genera that are recognized as having survived from the Mesozoic into the Genozoic. Of the seven Genozoic genera, four surxdve in the Recent fauna. According to the literature surveyed, there is no chrono- logical overlap between the Mesozoic genera and the Genozoic genera.

As noted by a number of researchers (e.g., Hickman, 1976: 1094; Szabô, 1980: 49; Das, 2002: 99) fossU pleurotomariids are difficult to classify objectively, since the criteria upon which fossil pleurotomariid genera are de- nned differ conspicuously from those applied to Geno- zoic genera. The monophyly and phylogenetic relationships of the surviving Genozoic genera have been con- ñrmed using molecular data from living representatives (e.g., Harasewych et al, 1997; Harasewych, 2002). By contrast, the relationships of Mesozoic genera and the species assigned to them are far less certain, as generic classification tends to be based on relatively few conspicuous morphological features (Table 1) especially those that are most easily derived from poorly preserved specimens and external or internal molds, while other characters are unconstrained and may vary widely. Szabô (1980: 49) commented that "almost all genera can be identified with certainty" on the basis of the shape of the whorl section and the surface of the whorls, as well as the position and width of the selenizone. Gonti and Szabô (1987: 43) raised a question as to the significance of the presence or absence of an umbilicus in pleurotomariid

THE NAUTILUS, Vol. 121, No. 2

MESOZOIC CENOZOIC

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GENERA OF PLEUROTOMARIIDAE

Mamoeatomaria Begg and Grant-Mackie, 2003

Tahua Begg and Grant-Mackie, 2003

Murihikua Begg and Grant-Mackie, 2003

Pleurotomaria Def ranee, 1826

Omatospira Pan, 1982

SfuoreJ/a KittI, 1891

Talantodiscus P. Fischer, 1885

Pyrgotrochus P. Fischer, 1885

Anoöotomaria SzBbó, 1980

Bathroíomaria Cox, 1956

Cychstomana Szabó, 1980

Laevitomaria Conti and Szabó, 1987

Oöome/Za Cox, 1959

Conotomaria Cox, 1959

Leptomaria E. Eudes-Deslongchamps, 1864

Indomaria Das, 20Q2

Chelotia Bayie /n P. Fischer, 1885

Entemnotrochus P. Fischer, 1885

Pemirochus P. Fischer, 1885

Bayemtrochus Harasewych, 2002

Mikadotmchus Lindholm, 1927

Figure 3. Geological ranges of the genera and subgenera within the family Pleurotomariidae, arranged by first occurrence in the fossil record.

classification. The presence or absence of an umbilicus is sufficient to distinguish the most basal dichotomy among living Pleurotomariidae, yet this feature remains unconstrained and may vary widely within most Mesozoic genera and even within some "species" as they are currently diagnosed. It is therefore not surprising that fossil species are frequently reassigned from one genus to another (e.g., Pyrgotrochus to Laevitomaria Conti and Szabô, 1987: 46; Perotrochus to Leptomaria, see Das et al., 2005: 331), especially as more numerous and better preserved specimens become available.

SYSTEMATICS

Family Pleurotomariidae Swainson, 1840 Genus Obornella Cox, 1959 Ohomella Cox, 1959: 238.

Type Species: Pleurotomaria plicopunctata J. A. Eudes-Deslongchamps, 1849 (By original designation). Bajocian (Middle Jurassic) of France.

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rowly open umbilicus and a strongly convex base. Surface sculpture consists of closely spaced coUabral costellae (usually dominant) and spiral threads. The periphery is commonly crenate. The selenizone is narrow, smooth, often projecting onto the upper whorl face near the periphery. The labral slit is short.

Remarks: Ohornella is known from strata of Toarcian (Lower Jurassic) to Oxfordian (Upper Jurassic) age (183 Ma to 156 Ma). Greatest diversity has been documented from Europe (see Gründel, 2003; Hagele, 2003), and this genus has also been reported from northeastern Iran (Majidifard, 2003) and the Kutch region of western India (Jaitley et al, 2000; Das et al., 2005).

Ohornella texta (Delpey, 1948) (Figure 4, reproduced from Delpey, 1948: pi. 2, ñg. 1)

Leptomaria texta Delpey, 1948: 9, pi. 2, fig. 1.

Original Description (Translated): "(Height: 25 mm; diameter 31.5 mm; number of whorls: 5). The strip [selenizone] is anterior, wide relative to the last turn, and develops/changes normally until the multicarinate stage, becoming a little convex. The sculpture is latticed. An umbilicus obscured by a lamina pierces the convex base.

It is similar to Fleurotomaria eudora d'Orbigny, 1850, but the selenizone of this Oxfordian species is concave between two carinae, which is noticeably different from the Malagasy form. Argovian of Ankirijy (coll. Hourcq)."

Remarks: Unfortunately, Delpey did not specify were her type material was deposited. Inquiries at the Muséum national d'Histoire naturelle in Paris as well as the Université de Paris revealed that the specimens are not deposited in their collections. The description is minimal, and the illustration of the single, partial specimen (Figure 4) is poor, showing a specimen in which the selenizone runs along the shell periphery, with the suture

1 cm Figure 4. Obomella texta (Delpey, 1948). Reproduction of original illustration (Delpey, 1948: pi. 2, fig. 1, as Leptomaria texta).

adpressed along the lower edge of the selenizone. These features preclude the inclusion of this species in Lep- tomaria E. Eudes-Deslongchamps, 1864, which is characterized by convex whorls, with the selenizone at mid- whorl. It is more likely that this taxon is referable to the genus Ohornella, in which the selenizone is situated closer to the periphery. Delpey's illustration resembles the partial specimen (Jaitly et al. 2000: 39, pi. 2, figs. 8A, B) from the contemporary Dhosa Oolite Member of the Kutsch region of India that was identified as Ohornella aff. granulata (J. Sowerby, 1818). Delpey's taxon is here transferred to Ohornella, but its generic affinities remain obscure until the type, which is the only known specimen, is located.

Ohornella thompsonorum new species. (Figures 5-12)

Description: Shell (Figures 5-12) small (holotype maximum diameter 42.2 mm, minimum diameter 34.6 mm, height 26.4 mm) low, turbiniform, consisting of approximately 7 teleoconch whorls. Base moderately convex, with narrow umbilicus (Figure 7, u). Spire angle 103-106°. Spire slightly convex in profile. Suture adpressed, joining pre\dous whorl at or just below peripheral bulge (Figure 6, pb). Protoconch and first 3-5 teleoconch whorls eroded, only final 2-4 teleoconch whorls well preserved. Weak shoulder present on whorls 3-5, becoming convex, rounded in final two whorls. Axial sculpture of oblique radial costae most pronounced on whorls 3-5 (40-50 per whorl), weaker on subsequent whorls (80-108 per whorl), producing cancellate gran- ules at intersections with spiral cords, especially at peripheral bulge and on either side of selenizone (Figure 10, sz). Number of strong, simple spiral cords 7-9 between suture and selenizone, 0-3 on selenizone, 2-4 between selenizone and peripheral bulge, 21-22 along base, between peripheral bulge and broad parietal callus (Figure 7, pc). Selenizone (Figure 10, sz) narrow, convex, with 0-3 spiral cords, and numerous strong to weak lunulae, situated just above peripheral bulge. Aperture ovate, roughly perpendicular to coiling axis. Outer lip smooth, portion below slit offset from portion above slit by 39°. Slit narrow (~2.5 mm), extending posteriorly 62° from end of suture. Lip thickest in columellar and basal region, forming broad parietal callus that partly overlaps the umbilicus.

Type Locality: Zakaraha, near Toliara (also spelled Tulear), southwestern Madagascar. 6-7 m below surface on plateau cut by river.

Type Material: Holotype, USNM 534480; paratype 1, USNM 534481; paratype 2 USNM 534482, all from the type locality.

Age: Oxfordian (Upper Jurassic).

Etymology: This new species honors Jon and Beverly Thompson for their many contributions and years of service to The Bailey-Matthews Shell Museum in Sanibel, Florida.


Figures 5-11. Ohomella thompsonoi-uni new species. 5. Apical, 6. apertural, and 7. basal views of the holotype, USNM 534480. 8. Apical and 9. dorsal \iews of parat^pe 1, USNM 534481. 10. Details of sculpture between suture and periphery on last three dorsal whorls of paratype 1. 11. Apical \iew of paratype 2, USNM 534482. Abbre\iations: p, periphery; pb, peripheral bulge; pc, parietal callus; s, suture; si, posterior limit of slit; sz, selenizone; u, umbilicus.

Remarks: The assignment of this new species in the genus Obornella is provisional. The type species of Obornella, Pleurotomaria plicopunctata J. A. Eudes- Deslongchamps, 1849, from the Bajocian (Middle Juras- sic) of France and England, is near one end of a morphological spectrum (see Hagele, 2003: fig. 10) that is distinguished by a low, conical spire and conspicuous axial fluting along the shell periphery and base. The other end of this morphological spectrum is characterized by shells with a higher, stepped spire, a rounder

aperture, and spiral cords along the base, features remi- niscent of the genus Pleurotomaria. Obornella thompsonorum more closely resembles O. trapeza (Hudleston, 1895: pi. 40, figs. 5a, b) and the "elevated variety" of O. granulata (Sowerby, 1818) illustrated by Hudleston (1895: pi. 40, figs, la, b) as Pleurotomaria granulata var. coplata Deslongchamps, 1848, but differs in having a less pronounced, more rounded peripheral bulge, a spire that is stepped along intermediate whorls, and strongly beaded sculpture along both sides of the selenizone. Two


European species of Obomella had been reported from the Kutch fauna (Jaitly et al., 2000; Das, et al., 2005). Ohornella wuerttemhergensis (Sieberer, 1908) from Lower Jurassic [upper Bathonian] strata, is much flatter, lacks a parietal callus, and has a far broader umbilicus than O. thompsonorum. Specimens of Obomella granú- lala (Sowerby, 1818) from the Dhosa oolite member of the Chan Formation [Oxfordian] of Kutsch (Jaitly et al., 2000: pi. 1, flgs. 5-7) resemble O. thompsonorum in overall proportions, but have more convex whorls between suture and periphery, far weaker axial sculpture, a broader umbilicus, and lack the distinctive parietal callus. Ohornella thompsonorum may be distinguished from Obomella texta (Delpey, 1948), the only prewously described pleurotomariid from the Jurassic of Madagascar, by the position of the selenizone above rather than along the periphery, and by hawng more prevalent sculpture, especially adjacent to the selenizone.

The shell microstructures found in Obomella thompsonorum (Figure 12) are similar to those of pleurotomariids from the Carboniferous (Batten, 1972), Triassic (Bändel, 1991), Jurassic (B0ggild, 1930), Cretaceous (Kiel, 2006), and Becent (Harasewych, 2002), indicating that shell microstructure is a very conservative character in this group.

Genus Bathrotomaria Cox, 1956

Type Species: Trochus reticulatus J. Sowerby, 1821. (By original designation). Kimmeridgian (Upper Juras- sic) of England.

Diagnosis: Bathrotomaria can be distinguished by its usually large (to 130 mm), trochiform shell with a spire that may be elevated to depressed. The umbilicus may be broad to entirely absent. The whorl profile is usually angulate and non-tuberculate, with a broad ramp and a second carina or angulation, just overlapped on the spire.

Figure 12. SEM image of a fracture surface of the shell of Obomella thompsonorum new species at the aperture just below the slit, showing the transition from the simple prismatic outer layer to the nacreous inner layer. The arrows indicate areas with recr^'stalized shell material. Abbreviations: Nac, na- cre; spr, simple prismatic crystals.

The selenizone is situated below the ramp angle. Surface sculpture of spiral cords and threads, commonly cancellate at intersection with coUabral threads. The selenizone is moderately broad, the labral slit short.

Remarks: The oldest member of this genus is Bathro- tomaria paipotensis Griindel, 2001, from the Sinemurian (Lower Jurassic) of northern Chile. The selenizone is narrow for the genus and the spiral sculpture is faint compared to other species. During the Middle and Up- per Jurassic the genus was diverse and widely distributed from Peru (Cox, 1956) to Europe, India, and the Afro- Arabian East coast (Cox, 1960, 1965; Howarth and Nor- ris, 1998; Das et ah, 2005), and possibly also New Zea- land (Gardner and Campbell, 1997), although the New Zealand record was not figured and needs confirmation. A number of species were reported from the Lower Cre- taceous of the Tethyan realm, including the largest species with 130 mm diameter (Das, 2002; KoUmann, 1982, 2002). From the Cenomanian (lower Upper Cretaceous) Kiel and Bändel (2004) reported four species of Bathro- tomaria from an intertidal rocky shore setting in Ger- many, the highest diversity at any Cretaceous locality. Further Upper Cretaceous records are few; the last record is from the lower Maastrichtian of France (Koll- mann and Odin, 2001).

Bathrotomaria annejoffeae new species (Figures 13-16)

Description: Shell (Figures 13-16) small for genus (holotype maximum diameter 47.7 mm, minimum diameter 43.8 mm, height 50.2 mm), with a tall, conical, strongly gradate spire, consisting of Q+ teleoconch whorls. Base weakly but evenly convex, with very narrow umbilicus (Figure 15, u). Spire angle 72°. Spire very slightly convex in profile. Suture weakly adpressed, joining previous whorl at or above peripheral bulge (Figure 14, pb). Protoconch and approximately first 2 teleoconch whorls missing. Subsequent early whorls with straight to weakly convex ramp between suture and ramp angle (Figure 14, ra) that becomes more inflated, convex on body whorl. Shell surface with broad, uneven, undulating rugae most evident near the suture. Axial sculpture of evenly spaced, weak axial costae (about 120 on body whorl) that produce weakly cancellate sculpture at intersections with spiral cords. Spiral sculpture (Figure 16) dominant, of 10-12 narrow, nearly abutting spiral cords between suture and ramp angle (comprised of single, smooth broad cord), 0-2 spiral cords between ramp angle and selenizone, 6-7 along selenizone, 3-4 between selenizone and peripheral bulge. Base with 36-38 spiral cords that are twice as broad as intervening spaces. Se- lenizone (Figure 16, sz) broad, weakly convex, nearly abutting ramp angle, spanning slightly more than half the distance between ramp angle and peripheral bulge. Ap- erture elongate, roughly pentagonal, long axis nearly perpendicular to the coiling axis. Outer lip broken. Slit broad (~3.8 mm), extending posteriorly 117° from the end of the suture. Lip thick along columellar region, with

M. G. Harasewych and S. Kiel, 2007 3

2 cm

Figures 13-16. Bathrotomaria annejojfeae new species. 13. Apical, 14. apertural, and 15. basal views of the holot^pe, USNM 534483. 16. Details of sculpture between suture and periphery on last three dorsal whorls of the holotype. Abbreviations: p, periphery; pb, peripheral bulge; pc, parietal callus; ra, ramp angle; s, suture; si, posterior limit of slit; sz, selenizone; u, umbilicus.

narrow, weakly reflected parietal fold that forms a narrow parietal callus that partially occludes the umbilicus.

Type Locality: Zakaraha, near Toliara, southwestern Madagascar, 6-7 m below surface on plateau cut by river.

Type Material: Holotype, USNM 534483, from the type locality.


Etymology: This new species honors Anne Joffe in

recognition of her many contributions and long service to the field of malacology, particularly to the American Ma- lacological Union (now American Malacological Society) and most recently to The Bailey-Matthews Shell Mu- seum in Sanibel, Florida.

Remarks: Bathrotomaria was among the most wide- spread and diverse of the pleurotomariid genera throughout the Middle and Upper Jurassic and Creta- ceous. This is particularly true of the Jurassic fauna of the


Kutch region, from which ten species have been reported (Maithani, 1967; Jaitly et al., 2000; Das et al, 2005). Of these, five are of Oxfordian age. From its contemporary congeners within the Indo-Madagascan province, Bathrotomaria annejoffeae is readily distinguished from B. tewarii (Maithani, 1967), B. huddhai Das et al., 2005, B. prasantai Das et al., 2005, and B. dhosaensis Das et al., 2005 in having a proportionally taller, more gradate spire and a narrower umbilicus. In shell profile, B. annejoffeae more closely resembles specimens of B. reticulata (Sowerby, 1821) (Das et al., 2005: figs. 4, A-G) and B. siebereri (Jaitly et al., 2000:pl. 3, ñgs. 2-3) both from the Bathonian of Kutsch, and the Oxfordian B. millepunctata (Eudes-Deslongchamps, 1849) (Jaitly et al., 2000: pi. 3, fig. 4), but lacks granular whorl angula- tions of B. siebereri and B. reticulata (see Das et al., 2005:334). Das et al. (2005: 334) also noted that the Kutch specimens of B. reticulata lack an umbilicus (present in B. annejoffeae), whereas the suture of B. siebereri is deeply canaliculated, unlike that of B. annejoffeae. Bathrotomaria annejoffeae can also be recognized on the basis of the columellar portion of the aperture being long, straight, and nearly co-axial.

Bathrotomaria bedetteae new species. (Figures 17-20)

Description: Shell (Figures 17-20) small for genus (holotype maximum diameter 65.0 mm, minimum diameter 58.8 mm, height 47.5 mm), with a short, broad, weakly gradate spire, consisting of 5+ teleoconch whorls. Base weakly but evenly convex, lacking an umbilicus (Figure 19). Spire angle 92°. Spire weakly convex in profile. Suture weakly adpressed, joining previous whorl at or above peripheral bulge (Figure 18, pb). Protoconch and approximately first 3 teleoconch whorls missing. Subsequent early whorls gradate, with straight ramp between suture and ramp angle (Figure 18, ra) that becomes more inflated, convex with increasing whorl number. Axial sculpture of broad, low, closely spaced, axial costae (about 56 on body whorl) that produce a coarsely cancellate sculpture at intersections of spiral cords, including weakly beaded ramp angle and peripheral bulge. Spiral sculpture codominant, of 3-A low, broad, cords between suture and ramp angle, 0-1 spiral cords between ramp angle and selenizone, 0-1 between selenizone and peripheral bulge. Base with 17-19 finer spiral cords that are 2-4 times as broad as intervening spaces. Selenizone (Figure 20, sz) broad, weakly convex, nearly abutting peripheral bulge, spanning slightly more than half the distance between ramp angle and peripheral bulge. Surface without spiral cords, sculpture limited to strong lunulae. Aperture elongate, weakly pentagonal, long axis deflected from the coiling axis by 103°. Outer lip damaged, slit morphology not known. Inner lip thickest along columellar region.




Etymology: This new species honors the late Barbara A. Bedette, whose 52 years of ser^dce to moUuscan paleontology at the National Museum of Natural History has benefited a multitude of researchers.

Remarks: Bathrotomaria bedetteae may be readily distinguished from B. annejoffeae, with which it co- occurs, on the basis of its lower, broader shell, with a weaker, more rounded ramp angle, by its coarser and more prominent cancellate sculpture that extends onto the ramp angle and peripheral bulge, and by the absence of an umbilicus. Bathrotomaria bedetteae is similar in profile to B. tewarii, B. prasantai, and B. dhosaensis, all from contemporary strata in Kutch, but differs in hawng a more gradate spire and a pronounced peripheral band, and in lacking an umbilicus.

Genus Leptomaria E. Eudes-Deslongchamps, 1864.

Type Species: Pleurotomaria amoena J. A. Eudes- Deslongchamps, 1849. (By original designation). Bajo- cian (Middle Jurassic) of France.

Diagnosis: Species of Leptomaria can be recognized on the basis of their large, turbiniform shells with low to moderately high spires and weakly to strongly rounded whorls. The umbilicus may range from broad to entirely absent. The whorl profile is rounded, lacking an angulate shoulder. The selenizone is situated at mid-whorl. Sur- face sculpture consists primarily of narrow spiral threads with finer axial threads forming weakly cancellate sculpture in some species.

Remarks: Leptomaria has been reported from strata ranging in age from Bajocian (Middle Jurassic, Knight et al., 1960) to Selandian (Paleocene, KoUmann and Peel, 1983). Hickman (1976) suggested that even some Eocene species may be included. Not surprisingly, the genus had a cosmopolitan distribution, with diverse faunas ranging from England (Cox, 1960) to New Zealand (Hudson, 2003). Leptomaria has been previously reported from Gretaceous deposits of NW Madagascar (Delpey, 1948; CoUignon, 1949; Kiel, 2006). It is represented in the Jurassic fauna of Kutch (Jaitly et al., 2000; Das et al., 2005) but is not as diverse as the genera Bathrotomaria or Pleurotomaria.

Leptomaria takahashii new species (Figures 21-24)

Description: Shell (Figures 21-24) moderately large for the genus (holotype maximum diameter 82.4 mm, minimum diameter 67.23 mm, height 57.3 mm) low, turbiniform, consisting of approximately 6 teleoconch whorls. Base broadly and evenly convex, with very narrow umbilicus (Figure 23, u). Spire angle 109°. Spire strongly convex in profile. Suture (Figure 24, s) abutting.


2 cm

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2 cm

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î^'îi-^' ,.Kî-

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Figures 17-20. Bathrotomaria hedetteae new species. 17. Apical, 18. apertural, and 19. basal views of the liolot)pe, USNM 534484. 20. Details of sculpture between suture and periphery on last three dorsal whorls of the holotype. Abbreviations: p, periphery; pb, peripheral bulge; ra, ramp angle; s, suture; si, posterior limit of slit; sz, selenizone.

joining previous whorl just below selenizone (Figure 24, sz) in early whorls, at or below periphery (Figure 22, 24, p) in later whorls. Protoconch and part of first teleoconch whorl eroded. Early teleoconch whorls (whorls 2-4) evenly rounded, later whorls (whorls 5-6) becoming more gradate, but evenly rounded, lacking a angular shoulder. Axial sculpture of numerous ñne growth striae that produce a strongly reticulate pattern most ewdent between adjacent cords in region between suture and selenizone of early whorls, and broader low, axial costae that form a weakly cancellate pattern at intersections with the spiral cords. Spiral sculpture dominant, with 10

strong, closely spaced cords between suture and selenizone, 7-9 cords between selenizone and periphery, and 29-32 cords along base. Spiral cords may become broader and less distinct with increasing whorl number. Selenizone (Figure 24, sz) narrow, convex, situated just above periphery, with a single, median spiral cord present in early whorls, absent in later whorls, and numerous strong to weak lunulae throughout its length. Aperture evenly ovate, long axis forming an angle of 103° with coiling axis. Outer lip smooth, portion below slit offset from portion above slit by 30°. Slit narrow (~3.3 mm), extending posteriorly 81° from end of suture. Lip


2 cm

21

22

24

Hsz

Figures 21-24. Leptomaria takahashii new species. 21. Apical, 22. apertural, and 23. basal \iews of the holotype, USNM 534485. 24. Details of sculpture between suture and periphery' on last three dorsal whorls of the holotype. Abbreviations: p, periphery; pc, parietal callus; s, suture; si, posterior limit of slit; sz, selenizone; u, umbilicus.

thickest in columellar and basal region, forming narrow parietal callus that partly overlaps the umbilicus.




Etymology: This new species honors Mr. Chris Ta- kahashi of Honolulu, Hawaii, in recognition of his many contributions to the study of li\dng and fossil moUusks.

Remarks: Leptomaria daityai Das et al., 2005 (Cal- lo\dan to Oxfordian of Kutch, India) occurs in coeval deposits, and most closely resembles L. takahashii in terms of size and general profile, but L. takahashii has a shorter spire, more rounded whorls, and has a very narrow rather than a wide umbilicus. Although the genus Leptomaria is well represented in the Jurassic fauna of the Indo- Madagascan region, most published records are either from Bathonian and Callowan strata of Kutch, or from Cretaceous deposits of Madagascar. Older taxa, including L. fraga (J. A. Eudes-Deslongchamps, 1849), L. sim- plex Jaitly et al., 2000, L. pseudoumbilicata Jaitly et al..


2000 (transferred to Bathrotomaria by Das et al., 2005: 340) are easily distinguished as they are smaller, and have a tall, conical whorl profile. Although the Bathonian L. asurai Das et al., 2005, is of comparable size, it differs in also hawng a tall conical profile and lacks an umbilicus.

DISCUSSION

The species most similar to those described herein occur in contemporaneous fossil deposits of northwestern In- dia. This is not surprising considering that the Madagas- car species lived near the southern tip of a long and narrow embayment between East and West Gondwana, while northwestern Indian species lived at the northern tip of this embayment, where it opened to the Tethys Ocean (Figure 2). Although Recent pleurotomariids are restricted to hard substrates along the outermost continental shelf and upper continental slope, Mesozoic pleurotomariids inhabited sublittoral depths along the continental shelf. Studies of Upper Jurassic phytogeography have indicated that southern Madagascar was part of a warm temperate biome, while northern Madagascar and northwestern India were part of a warmer, subtropical biome (Rees et al., 2000: fig. 7C). Based on analyses of stable isotope compositions of the ammonite Perisphinc- tes (Dichotomoceras), Lécuyer and Bûcher (2006: 7, fig. 3) reported seawater temperatures ranging from 21.5°G to 24.1°G in the Morondava Basin of southwestern Madagascar, during the Oxfordian stage of the Upper Jurassic.

Extensive studies of Jurassic pleurotomariid faunas within this Indo-Madagascan Province (e.g., Gox, 1965; Jaitly et al., 2000; Das, 2005) have documented the presence of the genera Bathrotomaria, Leptomaria, Obomella, Fleurotomaria, Anodomaria, and Pyrgotro- chus, the ñrst three by far the most diverse. As additional specimens from the Jurassic of Madagascar become available, it is likely that the generic composition and diversity of this fauna will mirror that of the Kutch region of northwestern India.

ACKNOWLEDGMENTS

We are grateful to Chris Takahashi for generously mak- ing the specimens available for study, and allowing the type material to be deposited in the collections of the National Museum of Natural History, Smithsonian Insti- tution. We thank K. Bändel, J. Hartmann, and W. Weitschat, (all Hamburg) and H. Keupp (Berlin), for their help regarding the localities and their age; and T. A. White for providing the Jurassic paleo-map. A. Oleinik and an anonymous reviewer are acknowledged for their

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Appendix 1. Supraspecific taxa included in the family Pleurotomariidae, listed in the order in which they were proposed. Their type species are provided, as are known stratigraphie ranges, and geographical distributions.

Pleurotomaria Defrance, 1826 - Type species, Trochus angli- cus J. Sowerby, 1818. Middle Triassic (Anisian; range extended by Begg and Grant-Mackie, 2003: 259) to Lower Cretaceous (Aptian). Cosmopolitan.

Leptomaria E. Eudes-Deslongchamps, 1864 - Type species, Pleurotomaria amoena J. A. Eudes-Deslongchamps, 1849. Middle Jurassic (Bajocian) to Paleocene (Danian). Cosmo- politan.

Talantodiscus Fischer, 1885 - Type species, Pleurotomaria mirabilis Deslongchamps, 1848. Middle Triassic (Kaihikuan ; range extended by Begg and Grant-Maclde, 2003: 259) to Middle Jurassic (Bajocian). Europe and New Zealand. Knight et al. (1960: 1214) included this genus in the family Porcelliidae Broili, 1924. Subsequent authors (e.g., Szabô, 1980: fig. 3; Begg and Grant-Mackie, 2003: 229) included this genus within Pleurotomariidae.

Pyrgotrochus P. Fischer, 1885 - T)pe species, Pleurotomaria hitorquata J. A. Eudes-Deslongchamps, 1849. Lower


Jurassic to Upper Cretaceous (Cenomanian). Cosmopoli- tan.

Perotrochus P. Fischer, 1885 - Type species, Pleurotomaria quoyana Fischer and Bernardi, 1856. Oligocène to Re- cent. Cosmopolitan.

Chelotia Bayle in P. Fischer, 1885 - Type species, Pleuroto- maria concava Deshayes, 1832. Eocene. Europe.

Entemnotrochus P. Fischer, 1885 - T)pe species, Pleuroto- maria adansoniana Crosse and Fischer, 1861. Eocene to Recent. Cosmopolitan.

Stuorella Kittl, 1891 - Type species, Trochus subconcavus Münster, 1841. Middle Triassic (Ladinian) to Upper Cre- taceous (Campanian; range extended by Kiel and Bändel, 2000). Europe.

Mikadotrochus Lindholm, 1927 - T)'pe species, Pleurotomaria beyrichii Hilgendorf, 1877. Western Pacific, Pliocene to Recent.

Bathrotomaria Cox, 1956 - Type species, Trochus reticulatus J. Sowerby, 1821, by original designation. Lower Jurassic to Upper Cretaceous (Senonian). Cosmopolitan.

Conotomaria Cox, 1959 - T)pe species, Pleurotomaria mail- leana d'Orbigny, 1843. Middle Jurassic (Bajocian) to Pa- leoceno (Danian; range extended by Pacaud, 2004). Cos- mopolitan.

Ohomella Cox, 1959 - Type species, Pleurotomaria plicopunctata J. A. Eudes-Deslongchamps, 1849. Lower Jurassic (Toarcian) to Upper Jurassic (Oxfordian) Cosmopolitan.

Anodotomaria Szabô, 1980 [described as a subgenus of Pleu- rotomaria] - Type species Pleurotomaria scacchi Gem- mellaro, 1874. Lower Jurassic (Pliensbachian) to Middle

Jurassic (Callovian; range extended by Jaitly et al., 2000: 36). Europe to northwestern India.

Cyclostomaria Szabô, 1980 - Type species, Pleurotomaria suessi Homes, 1853. Lower Jurassic (Pliensbachian) to Middle Jurassic. Europe.

Ornatospira Pan, 1982 - Type species Omatospira mirifira Pan, 1982. Triassic, China.

Laevitomaria Conti and Szabô, 1987 - Type species, Pyrgotro- chus? problematicus Szabô, 1980. Lower Jurassic (Pliens- bachian) to Middle Jurassic (Bajocian) Europe. Question- ably included in Pleurotomariidae by the authors.

Indomaria Das, 2002 [described as a subgenus of Pleuroto- maria] • Type species, Pleurotomaria (Indomaria) umien- sis Das, 2002. Lower Cretaceous (Berriasian). Northwest- ern India.

Bayerotrochus Harasewych, 2002 - Type species, Pleuroto- maria midas Bayer, 1964. Miocene to Recent. Cosmopoli- tan.

Murihikua Begg and Grant-Mackie, 2003 - Type species, Murihikua tuhawaiki Begg and Grant-Mackie, 2003, Middle Triassic (Aniasian) to Upper Triassic (Norian). New Zealand.

Tahua Begg and Grant-Mackie, 2003 - Type species. Tahua waipiro Begg and Grant-Mackie, 2003 Middle Triassic (Aniasian) to Late Triassic (Norian). New Zealand.

Mamoeatomaria Begg and Grant-Mackie, 2006 (new name for Mamoea Begg and Grant-Mackie, 2003) - Type species, Mamoea wairakiensis Begg and Grant-Mackie, 2003. Middle Triassic (Aniasian to Ladinian). New Zea- land.

upper jurassic pleurotomariidae (gastropoda) from … · 2019. 12. 14. · jurassic) age....

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